Anales del Jardín Botánico de Madrid 61(2): 155-180 www.rjb.csic.es Check-list of the Basque coast benthic algae (North of Spain) por José M. Gorostiaga1, Alberto Santolaria1, Antonio Secilla1, Coro Casares2 & Isabel Díez1 1 Departamento de Biología Vegetal y Ecología, Facultad de Ciencias, Universidad del País Vasco, Apdo. 644, E-48080 Bilbao, Spain [email protected] (corresponding author) [email protected] [email protected] [email protected] 2 I.E.S. Usandizaga, Av. de Felipe IV, E-20011 San Sebastián, Spain. [email protected] Abstract Resumen An annotated check-list of the marine benthic flora of the Span- Se presenta un catálogo de la flora bentónica marina de la costa ish Basque coast, based on literature records and original data, vasco-española basado en citas bibliográficas y algunos datos is presented. According to the present taxonomy, the known list originales. De acuerdo con la taxonomía actual, la lista de algas of algae totals 372 taxa (354 species and 18 infraspecific taxa), conocidas asciende a 372 táxones (354 especies y 18 táxones in- distributed as follows: 23 Cyanophyta, 221 Rhodophyta, 74 fraespecíficos), que se distribuyen como sigue: 23 Cyanophyta, Heterokontophyta and 54 Chlorophyta. In addition, 16 taxa are 221 Rhodophyta, 74 Heterokontophyta y 54 Chlorophyta. considered as taxa inquirenda and 8 as taxa excludenda. For Además, 16 táxones se consideran como taxa inquirenda y 8 each taxon the vertical distribution, abundance estimation, and como taxa excludenda. Para cada taxon se ofrece información two first published records are given. Remarks on the most note- sobre su distribución vertical, una estima de su abundancia y las worthy features of the flora of the study area are included. The dos primeras citas publicadas. Se incluyen comentarios sobre las floristic character of Spanish Basque coast flora was compared características más destacables de la flora del área de estudio. Se with nearby regions applying Cheney’s ratio [(Rhodophy- compara el carácter florístico de la flora de la costa vasco-es- ta+Chlorophyta)/Phaeophyta, or (R+C)/P]. The resulting high pañola con respecto a las regiones próximas aplicando el índice value (4,09) indicates its warm-water character, which was al- de Cheney [(Rhodophyta+Chlorophyta)/Phaeophyta, o (R+C)/P]. ready noted at the end of the XIX century. From a physionomical El elevado valor resultante (4,09) indica su carácter meridional, point of view, Basque benthic vegetation resembles more close- lo que ha sido señalado desde finales del siglo XIX. Desde un pun- ly that of southern regions (S Portugal, Morocco). The warming to de vista fisionómico la vegetación bentónica vasca se aseme- process of waters off the Basque coast during summer up to ja más a la de regiones meridionales (sur de Portugal, Marrue- 22 ºC is responsible for the absence or very rare occurrence of cos). El calentamiento de las aguas de la costa vasca hasta 22 ºC cold temperate species like large fucoids and kelps, which are durante el verano es responsable de la falta o rareza de especies common in Galicia and Brittany, and explains the abundance of templado-frías como las fucáceas y laminariales, comunes en numerous warm-temperate species as ceramiaceous algae. Galicia y Bretaña, y explica la abundancia de numerosas especies templado-cálidas como las ceramiáceas. Key words: benthic marine algae, biogeography, Cantabrian Palabras clave: algas bentónicas marinas, biogeografía, Sea, Chlorophyceae, Cyanophyceae, nomenclature, Phaeo- cianofíceas, clorofíceas, feofíceas, macroalgas, mar Cantábrico, phyceae, Rhodophyceae, seaweeds, taxonomy. nomenclatura, rodofíceas, taxonomía. Introduction Atlantic coast, such as England, Brittany, Galicia, Por- tugal and the nearby French Basque coast. The first list Information concerning the algal flora of the Span- of marine macroalgae including Spanish Basque coast ish Basque coast has been sparse and late-coming com- records was compiled by Colmeiro (1867) who report- pared to information on other regions of the European ed 12 new records of Rhodophyta, 8 Heterokontophy- 156 Anales del Jardín Botánico de Madrid 61(2) 2004 ta and 4 Chlorophyta. Several works, mainly of Fisher- Taxa are named with their nomenclatural authori- Piette (1935, 1955, 1963), increased the number of ties, and the names have been abbreviated following known species to 46 by the year 1965. The following Brummitt & Powell (1992). In the Cyanophyta and noteworthy contribution was made by Hoek & Donze Chlorophyta orders and families are arranged accord- (1966), who reported 68 new records, including ing to Silva & al. (1996), with some remarks by Cyanophyta. This work was the main reference source Komárek & Anagnostidis (1986) and Anagnostidis & during the 1970’s, a decade with few algological contri- Komárek (1988), for the first case. In Rhodophyta, we butions. However, the knowledge of Basque seaweeds follow Garbary & Gabrielson (1990), with the addi- increased particularly during the 1980’s (Angulo, 1980; tion of the Ahnfeltiales (Maggs & Pueschel 1989), the Ibáñez & al., 1980; Gorostiaga, 1981; Gorostiaga & Li- Gracilariales (Fredericq & Hommersand 1989) and mia, 1985; Gorostiaga & al., 1987,1988; Casares, 1988, the Plocamiales (Saunders & Kraft 1994), while the 1989; Casares & Seoane-Camba, 1988, 1989), when the order Ceramiales is arranged according to Maggs number of known algae reached 270 taxa. Recent con- & Hommersand (1993). The systematic treatment of tributions have finally leveled the knowledge of the Phaeophyceae follows Wynne & Kraft (1981). All Spanish Basque coast algal flora with nearby areas genera and species are arranged alphabetically. (Gorostiaga & Santolaria, 1992; Bárbara & al., 1995; The present taxonomic status and nomenclatural Gorostiaga, 1995; Díez & al., 1996; Secilla & al., 1997, changes of the taxa concerned have been considered. 2000; Gorostiaga & al., 1998; Santolaria & al., 1998). The check-list includes the synonyms under which a The aim of the present work is to compile for the specific or infraspecific taxon has been recorded. first time the marine algal flora (Cyanophyta, Rhodo- Records that need to be confirmed are included in a phyta, Heterokontophyta and Chlorophyta) of the separate list as taxa inquirenda. Likewise, a list of taxa Spanish Basque coast. For this purpose a critical re- excludenda is also given with pertinent comments and view of all published records has been carried out and references. Specimens of most of the species included we have supplemented this information with 37 new in this check-list are deposited in the herbaria of the records of our collections. This check-list contributes Basque Country University (BIO), the University of to increasing the understanding of the biogeographi- Barcelona (BCF) and the University of Santiago de cal characteristics of this Atlantic area and the knowl- Compostela (SANT). Information about the abun- edge about its biodiversity. dance estimation and vertical distribution of each species is given together with an indication of the Material and Methods provinces (Gipuzkoa and Bizkaia) from which they The Spanish Basque coast is located in the northern have been reported. part of the Iberian Peninsula, on the south-eastern cor- The floristic character of Spanish Basque coast flora in ner of the Bay of Biscay (Fig. 1). It is mostly rocky with relation to cold-water and warm-water poles was com- an exposed to very exposed shore and very few estuar- pared with that of southern England (Hardy & Guiry, ies, all of them small. Following the coastline and in- 2003), Brittany (Feldmann, 1954; Feldmann & Magne, cluding the estuaries, it extends over 300 km from Cape 1964; Coppejans, 1995), Galicia (Miranda, 1934; Donze, Higer (43º23’46’’N; 1º47’24’’W) in the East to Kobaron 1968; Pérez-Cirera, 1975a, 1975b; Niell, 1978; López Point (43º21’01” N; 3º09’58” W) in the West. In polit- Rodríguez & Pérez-Cirera,1990, 1995, 1996; López Ro- ical terms it comprises the Spanish provinces of Gi- dríguez & al., 1991, 2003; Gallardo & Margalet, 1992; puzkoa and Bizkaia. The sea surface temperature Granja & al., 1992, 1993; Guillermes & Cremades, 1993, ranges from 11 ºC to 22 ºC in February and August, re- 1997; Guillermes & al., 1994; Bárbara & Cremades, spectively (Ibáñez, 1990; Borja & al., 2000). Biogeo- 1996; Otero-Schmitt & Pérez-Cirera, 1996, 1998, 2002; graphically, it belongs to the warm temperate NE At- Cremades & al., 1997b, 2002; Veiga & al., 1997, 1998a, lantic sub-region (Hoek & Breeman, 1990). 1998b; Bárbara & al., 2002, 2003; Calvo & Bárbara, The list of marine benthic algae (Cyanophyta, 2002, 2003, in press; Peña & Bárbara, 2002, 2003a, Rhodophyta, Heterokontophyta and Chlorophyta) 2003b), the Atlantic coast of the southern Iberian Penin- was compiled after a careful review of all published sula (Ardré, 1970, 1971; Flores Moya & al., 1995a, records and supplemented by our own collections. 1995b; Conde & al., 1996; Conde & Flores Moya, 2000; Unpublished works such as M. Phil. Thesis, Ph.D. Báez & al., 2001), Morocco (Benhissoune & al., 2001, Doctoral Thesis and Technical Reports have not been 2002a, 2002b, 2003), the Canary Islands (Haroun & al., considered. The data presented here date from Mon- 2002) and the Mediterranean coast of Andalusia (Flores tagne (1842) to Díez & al. (2003). First and second Moya & al., 1995a, 1995b; Conde & al., 1996) using records are given for each taxon. Cheney’s ratio [(R+C)/P] (Cheney, 1977). J.M. Gorostiaga & al.: Check-list of the Basque coast benthic